Beruflich Dokumente
Kultur Dokumente
^ 243
Ecophysiology of xerophytic and halophytic
vegetation of a coastal alluvial plain in
northern Venezuela
I. Site description and plant communities
BY E. MEDINA' , W, J. CRAM", H, S. J. LEE'^^ U. LUTTGE' ' *, M. POPP' ',
J. A, C. SMITH' ' AND M. DIAZ'
' Centro de Ecologia y Ciencias Ambientales, Instituto Venezolano de Investigaciones
Cientificas, Caracas 1020-A, Venezuela
'^Department of Biology, The University, Newcastle upon Tyne, NEl 7RU, UK
'^ Institut fur Botanik, Technische Hochschide Darmstadt, D-6100 Darmstadt, FRG
^ Institut fiir Pflanzenpliysiologie der Universitdt, A-1091 Wien, Austria
^ Institut fiir Angewandte Botanik, Westfdlische Wilhelms- Universitdt,
D-4400 Munster, FRG
*' Department of Botany, University of Edinburgh, Edinburgh, EH9 3JH, UK
' Centro de Investigaciones en Ecologia y Zonas Aridas, Universidad Naciottal
Experimental Erancisco de Miranda, Coro, Venezuela
{Received 8 March 1988; accepted 22 July 1988)
SUMMARY
This paper describes the ecology of a coastal alluvial plain at Chichiriviche in northern Venezuela. The area
supports a great diversity of plant communities, ranging from mangroves on the seaward edge of the plain to non-
halophytic, fresh-water communities on the landward side. Small differences on topography result in a mosaic of
saline and less-saline environments. Rainfall is strongly seasonal, causing superficial flooding of the alluvial plain
in the rainy season and the creation of a hypersaline substratum during the dry season. As a consequence, much
of the plain is devoid oi vegetation. Towards the landward side of the plain there are numerous small 'vegetation
islands', fringed by halophilic succulent herbs, and made up of deciduous and semi-deciduous shrubs and trees
together with non-halophytic CAMplants such as cacti and bromeliads. In subsequent papers the results of
ecophysiological studies of these diverse plant species are presented.
Key words: CAM plants; epiphytes; halophytes, mangroves; salt flat; Venezuela,
INTRODUCTION habitat patchiness. Deciduous and semi-deciduous
non-halophytic shrubs and trees coexist with man-
Salt flats associated with estuarine mangroves under groves and shrubby and herbaceous true halophytes,
strong seasonal rainfall in the tropics constitute an Within this heterogeneous vegetation terrestrial and
interface between terrestrial halophytic and non- epiphytic CAMplants may constitute a significant
halophytic vegetation. Proximity to the coast and percentage of the total biomass, as is the case of the
seasonal distribution of rainfall result in pronounced columnar cacti and opuntias, and Bromehaceae of
seasonal variation in soil sahnity and availabihty of the genera Tillandsia (epiphytic) and Bromelia
superficial fresh water. Under these conditions an (terrestrial). Coexistence of species such as succulent
array of higher plants with contrasting ecophy- halophytes, with high chloride concentrations in
siological properties coexist due to microtopo- their leaf sap, and succulent CAMplants, with
graphical heterogeneity resulting in a high degree of negligible chloride content, has been explained on
* To whomall correspondence should be addres,sed, the basis of microhabitat differentiation and dif-
234 E. Medina and others
1 CHI CHI RI VI CHE
2 TOCUYO DE LA COSTA
3 CURAMI CHATE
* STODY SI TE
Figure 1. Location of the study site on the northern coast ot Venezuela
ferential distribution of active absorbing roots
(Walter, 1973). Non-halophytic cacti and terrestrial
bromeliads have supeflcial root systems which are
active only during the rainy season, when superficial
soil salinity is washed away by free running water.
Shrubby and herbaceous halophytes on the contrary,
are able to obtain water and nutrients from deeper
soil layers, with higher chloride content throughout
the year. The present project aims to document the
variety of ecophysiological types co-occurring in such
an environment, and characterise differentiation of
microhabitats. The field work was undertaken
during the months of November-December 1985
and March-April 1986.
create a mosaic of saline and less or non-saline soil
environments. Areas in lower geomorphological
levels are heavily flooded during the rainy season but
become brackish and salty as the dry season
progresses. These areas may be completely devoid or
vegetation, probably as a result of the strong seasonal
changes in salinity and flooding (Walter, 1973;
Walter & Breckle, 1984). Towards the Chichiriviche
bay the influence of sea water becomes more
pronounced and flooding tends to be permanent.
There, a typical estuarinc mangrove vegetation
dominates the landscape. At the other extreme,
deciduous forest vegetation surrounds the areas
flooded seasonally by the Tocuyo river.
STl' DV ARK A
The study area, locally known as the Cienega el
Ostional, is located about 6 km west of the town of
Chichiriviche in Estado Falcon in north-western
Venezuela ( 1O55' N, 68 21' W) (Fig. 1). The
whole area is made up of alluvial sand plains and
harrier beaches created by the Tocuyo and Tucurerc
rivers (Goddard & Piccard, 1972). Beaches extend
for about 30 km from Chichiriviche in the direction
of Curamichate. These alluvial plains are period-
ically flooded by the Tocuyo river, exerting a
superfical salt washing effect. Saline influence arises
from upward percolation of sea water from the
Chichiriviche bay. Small differences in topography
CLIMATE
Long-term climatoiogical data are available from the
town of Tocuyo de la Costa, approximately 16 km
north-west of the study site (Fig. 1) for the period
1963-86. Annual rainfall averages 1029 mm. The
annual distribution of rainfall (Fig. 2) shows two
clear peaks, a small one in April and a very
pronounced one in November. There is no rainless
month, even in the driest years, and only the months
of February and March present average rainfall
helow 40 mm. This climate type is very favourable
for succulents (Walter, 1973), including epiphytes,
which have the possibility of recharging their \yater
reserves shortly after significant rains. Potential
Ecophysiologv of xerophvtic and halophvtic vegetation in Venezuela. I 235
E
d 200
o
Q.
a
o 100
o
o
CC
TOCUYO DE LA COSTA (20m)
(1963-1985) o Rai nf al l av 1020 mm
Evaporation av 2325 mm
A Temperature av 26 4 "C
O
o
30 S.
E
cu
2 0 ' "
10
J F M A M J J A S O N D
Figure 2. Axera^c tnonllily \alue.s of raitil'all, cxaporatioti
and temperature tiir a 22 years period as measured in the
meteorological station of the Ministerio del .Ambiente in
Tocuyo dc la Costa, Estado I'alcon, Venezuela.
evaporation is very high throughout the year and is
below the rainfall curve only during November and
December. The evaporation curve shows two small
peaks, the first in March and the second in August,
in both cases just before the onset of the respective
rainy periods, associated with the apparent path of
the sun towards and from higher latitudes during the
summer in the northern hemisphere. Lowest evap-
oration rates are registered in the period November
to February, when they reach 5-5+ 0-3 mm d^';
during the rest of the year daily evaporation is signifi-
cantly higher reaching 6-4 + 0-3 mmd"'. Average
monthly temperature varies very little during the
year; daily temperature differences between mean
maxima and mean minima (7-1 C on average) are
larger than the differences between mean tempera-
tures of the hottest and the coldest month (August
and January respectively), which amounts to 2-1 C.
The period analysed shows a pronounced rainfall
Variation between successive years (Fig. 3). This
variation is of ecological importance because it can
regulate the expansion or contraction of the halo-
phobic vegetation through the degree of surface-soil
Washing occurring every year. Besides, occurrence of
humid years probably causes erosive impacts which
rnay reduce the soil level in the vegetation covered
islands, thereby decreasing survival of non-halo-
phytic elements. The 23-year observation period can
be roughly diflerentiated into two phases. The
period between 1964 and 1975 was rather humid
vv'ith a rainfall average of 1170 348 mmy"'. The
period from 1976 to 1986 was much drier with a
rainfall average of 856 + 357 mm y"\ Out of 23
years, 7 years had rainfall above average plus twice
the standard error of the mean, and of those, 5
occurred before 1975. In contrast, 7 out of 9 years
with rainfall below average minus twice the standard
2000-
TOCUYO DE LA COSTA {20m)
64 68 72 76
Ye a r s
80 84
Figure 3. .\tintial variations in total rainfall registered
the last 2.^ \ears mthe TocuNO df la Costa station.
ISO
150-
70-
60-
50-
40-
_ 30
E
l 2 0
10
AL
15 20 25 30
October
5 10 15 20 25 30 5 10 15
November December 1985
15 20 25 28 5 10 15 20 25 30 5 10 15
February March April 1986
Figure 4. Daily rainfall distributioti during the study
period of the rainy seasoti ot 1985 and the dry season ot
1986.
error occurred after 1975. However, variability is
high. In the month of November 1985, during the
period when the first series of field measurements
were undertaken, rainfall was nearly twice the long-
term average recorded in Fig. 3. Rainfall figures
registered durmg the periods of fieldwork charac-
terize the strong seasonal distribution of rain (Fig. 4).
During November 1985, 8d had rainfall x'alues
above 25 mm, among them 1 d with more than
150 mm.
V II G K T A T I O N
The vegetation in the Cienega el Ostional is
diflerentiated between the extremes of mangroves
and deciduous forest. Between these two extremes a
mixture of halophylic and halophobic vegetation is
236 E. Medina and others
Figure 5. General vegetation map of the study area
developed from aerial photographs provided by the
Ministerio del Ambiente, Caracas. L, lagoon with brackish
water; M, tall mangrove vegetation; B, deciduous forest,
G, secondary grassland; I.V., island vegetation complex;
vf, mainly vegetation-free, flooding-prone areas; C, long
term fresh-water flooded areas dominated by sedges; S,
the broken line indicates the border of areas of pre-
dominantly saline soils in the east from higher non-flooded
areas in the west.
found (Fig. 5). The lowest geomorphological level is
occupied by a lagoon (L) with brackish water and is
completely surrounded by tall mangrove vegetation
(M) [mainly Rhizophora mangle L. and Avicennia
germinans (L.) Stearn and scattered Laguncularia
racemosa CJaertn.]. The highest geomorphological
level is occupied by several areas of deciduous forest
(B) and grassland (G), probably of secondary origin,
dominated by the grass, Paspalum plicatulum Michx.
The area between the mangroves and the deciduous
forests contains a mosaic of vegetation units desig-
nated by us as an island-vegetation complex (I.V.)
where very slight changes in the soil level, in the
order of 5-20 cm, determine changes in salinity and
flooding and the coexistence of halophylic and
halophobic plant species. Between the mangrove
area and the island-vegetation units extend areas
completely devoid of vegetation or occupied by
sparse pure stands of Batis maritima L. and/or
Sesuvium portulacastrum L. (vf). In some areas fresh
water accumulates probably as a result of impeded
drainage by the elevated road. These areas are
occupied by almost pure stands of a few species of
Cyperaceae (C).
Vegetation units emerging from the plain, ex-
cluding pure mangrove associations, can be dif-
ferentiated as follows:
(1) Flooded areas with fresh water accumulated
during the rainy season. These areas are dominated
by species of Cyperaceae such as Eleocharis geniculala
R. Br. Prod., E. nutans R. Br. Prod., Fymbristylis
cymosa Vahl and F. spadicea Vahl. Some species
such as Nyniphaea sp., with floating leaves in open
water, are ephemeral and disappear during the dry
season (Fig. be).
(2) A Batis maritima-Sesuvium protulacastrinn
unit, bordering the vegetation-free salt fiats, which
can be temporarily flooded after heavy rains [Fig.
6(a) and (6)].
(3) A grassland elevated between 5-10 cm from
the salt flat dominated by Sporobolus virginicus
Kunth, intermixed with Oxycarpha suaedifolia Blake
and isolated individuals of the cacti, Opuntia zven-
tiana Britton & Rose and Acanthocereus tetragonus
(L.) Humik. (Fig. 6a).
(4) Small islands, 3-10 m in diameter, where the
soil surface can be 10-40 cmhigher than the salt-fiat
[Fig. 6(c) and {d)~\. The vegetation on top of these
islands is frequently dominated by a mangrove
associated species, Conocarpus erectus L. (Tom-
linson, 1986), reaching approximately 2 min stature,
frequently accompanied by a columnar cactus Sub-
pilosocereus ottonis Backeb., and a broad-joined
Opuntia species (unit 4a in Table 1). On isiands
without C. erectus a number of isoiated non-
halophylic tree species such as Prosopis fuliflora
D.C., Capparis hastata Jacq. and Maytenus karstenit
Reiss. may be found. Another occasional species is
the cactus, Pereskia guamacho Weber, a small
deciduous tree or shrub covered with succulent
fleshy ieaves ciuring the rainy season (unit 4/; in
Tabie 1). As an epiphyte on the shrubs and cacti, the
bromeliad Tillandsia flexuosa Sw. is abundant. In
islands nearby and in deciduous lorest relicts, tiie
orchid Schomburgkia humboldtiana Reichb. is also
frequently found. The floor of these islands may be
covered by dense thickets of Bromelia humilis Jacq-
or scattered tufts of Sporobolus virginicus.
The contact between the islands and the sand
plains on lower ground is occupied by the Balis-'
Sesuvium community. During the rainy season the
reddish-flowered Portulaca riibricaulis H.B.K., with
well developed underground tubercules, is quite
often seen.
(5) The deciduous woodland. These areas are
represented hy deciduous forest remnants of variable
size which have been clearly isolated from the main
forest core by erosive influences of running water
during humid years, but also to some extent by
human infiuence (Fig. bf). The erosion reduces the
soil level thereby increasing salinity percolation from
underground brackish water in those areas con-
stituting the transition zone between saline and non-
Ecophysiology of xerophytic and halophytic vegetation in Venezuela. I 237
Figure 6. Vegetation units within the study area, {a) General view of the alluvial plains showing Sporobolus
virginicus and Acanthocereus tetragonus (foreground), Batis maritifna and Sesuvium portulacastrum (middle) and
Conocarpus erectus. Background shows the main core of deciduous forest towards the north-east border of the
study area, (6) Edge of the vegetation zone with Sesuvium potulacastrum leading to higher ground with mixed
grass species, (c) View of the island vegetation in the rainy season, (d) View of a typical island during the dry
season, (e) Example of fresh water vegetation with Nymphaea sp, and Eleocharis geniculata. (f) Remnants of
deciduous forest vegetation.
saline soils. The borders of these forested areas
^re characterised by the presence of Capparis
odoratissima Jacq,, C. hastata, Capparis cf. pachaca
Jacq. (broad, round leaves), Caesalpinia coriaria
Willd, Prosopis juliflora, Jacquinia revoluta Jacq,,
^aytenus karenii, Erythroxylon cumanense H, B, K,
3nd shrubs of Croton sp, and Pereskia guamacho.
Occasionally there are dense thickets of the terrestrial
Dromeliads B. humilis and B, chrysantha Jacq. As
Epiphytes the bromeliads Tillandsia flexuosa and T.
''ecurvata L,, and the orchids Schomburgkia hum-
"oldtiana and Brassovala nodosa Hook, are commonly
found.
A profile through the study area which includes all
the vegetation units described above and a number
of the most representative species shows the species
distribution pattern associated with topographical
position and the fluctuation of the water table
(Fig, 7),
SOI LS
It is clear that distribution of vegetation units is
determined by superficial soil salinity resulting from
underground water percolation, which varies sea-
sonally because of rain washing. During the rainy
238 E. Medina and others
metres 20 AO 60 80 100 120 UO 160 180 200 220 240 260 280
Bare soil
Conocarpus erectus
Cupparis flexuosa
Prosopis juliflora
Eugenia sp.
Croton heliaster
Capparis odoratissima
Mimosa oligocantha
Capparis hastata
Maytenus karstenii
Trichilia trifoliata
Guapira sp.
Capparis Iinearis
Fimbristylis spadicea
Fimbristylis cymosa
Carex sp.
Desmanthus virgatus
Paspalum pUcatulum
Leptochioa sp.
Sporobolus virginicus
Acanthocereus tetragonus
Opuntia wentiana
Subpiiosocereus ottonis
Bastardia viscosa
Cissus sicyoides
Anthurium crassinervium
Philodendron sp.
Evolvulus tenuis
Sesuvium portulacastrum
Batis maritima
Oxycarpha suaedifolia
Bromeiia humilis
Tillandsia flexuosa
Phoradendron mucronatur
Tillandsia recurvata
cm
150
100
50
island
deciduous forest
halophytes grassland remnant grassland halophytes
- soil pit
water table - rainy season
soil pit
water table -dry season
Figure 7. Transect indicating location of the three soil pits excavated for measurement of ground-water depth
and vertical variations in soil salinity. The transect reveals distribution of most frfqiicnt plant species as related
to topographical variations.
season differences in soil salinity (expressed as
chloride content) among vegetation units are re-
latively small but it is clear that salinity decreases
from the salt fiats towards the deciduous forest
remnants (Table 1). Differences among vegetation
units increase markedly during the dry season. In
April 1986, in the top 0-10 msoil, salinity decreases
from a high of 340 mequiv. Cr kg"' soil in the salt
flats to a low of 18 mequiv. Cr kg"' in tbe deciduous
forest remnants. The sodium/cbloride equivalent
ratio was O-88 + O-O3, indicating tbat most of tbe salt
in these soils is NaCI. Seasonal variation in salinity is
quite large in tbe salt flats, particulary in the
Batis-Sesuvium zone (unit 2) and the Sporobotus
zone (unit 3). Those are the typical halophylic plant
communities along the Caribbean coast in Ven-
ezuela. Least variation in salinity is observed in the
remnants of the deciduous forests (unit 5). In-
dividuals of Subpitosocereiis ottonis are found only in
tbese areas vvbcre the superficial soil does not
increase markedly in chloride concentration during
tbe dry season. However, Opuntia sp. often occur
within Sporobolus grasslands, therefore their roots
reach into the soil layer of high salt content during
the dry season. Individuals oi B. humilis are found in
a wider range of superficial soil salinity. For this
species however, soil chloride content is of no
consequence because functional roots are not in tbe
soil but grow between the leaf sheaths into the tank
constituted by tbe leaf bases (Pittendrigh, 1948).
Soil profiles were excavated during the dry season
(in March and April 1986 and 1987) to determine the
vertical variations in nutrients and salinity and to
measure tbe depth of the ground water (Fig. 8).
Sodium and chloride were by far tbe dominant ion
species in tbe profile. The ground water from the
tbree vegetation sites (units 1, 4a and 4-b) had
significantly higher concentrations than sea water
collected nearby (Fig. 9). In all cases Cl con-
centration increased witb soil depth (Fig. 10).
Moreover, there vvere considerable differences in the
profiles depending on the location in relation to the
vegetation units. Down to 0-50 m in a profile dug
near a 6-5 mtall Subpiiosocereus ottonis at the border
of a deciduous forest remnant (Soil pit II, Fig. 9),
salinity was about half of that measured in a profile
in the salt plain near a Conocarpus erectiis-covercd
island (Soil pit I). At lower levels, nearer the lagoon
Ecophysiology of xerophytic and halop/iytic vegetation in Venezuela. I
239
Table 1. Seasonal changes in superficial soil chloride contents. Cienega el Ostional Chichiriviche, Stado Falcon
(see text for description of vegetation units)
(1) Vegetation-free salt flats
(2) Batis + Scsuiiuni zone
(3) Spnrobolus + Oxxcarpha zone
4() Conocarpus island with
Bromelia humilis
4(6) Non-halophilic trees and
Bromelia sp. underneath, moss
covered soil surface
(5) Deciduous forest with
Bromelia sp. beneath
Chloride content
Depth (cni)
0-10
10-20
0-10
0-10
10 20
0-10
10-20
0-10
10-20
0-10
10-20
in nict|uiv. kj
Dry season
340+ 133
219 + 49
203 + 107
110 + 87
70 + 26
46 + 34
114+149
28 + 20
44 + 9
18 + 21
21+27
X ' dry soil
Rainy season
65
107
37
17
11-36
11
16